The destructive invasive pathogen Phytophthora lateralis found on Chamaecyparis lawsoniana across the UK

In 2010–2011, Phytophthora lateralis was isolated from diseased Chamaecyparis lawsoniana exhibiting dieback and mortality at eight geographically separate forest, parkland and shelterbelt locations in England, Scotland and Northern Ireland. In 2011, P. lateralis was also isolated from young symptomatic nursery plants of C. lawsoniana and Thuja occidentalis recently imported into Scotland from mainland Europe. These are the first findings of P. lateralis in the UK. At six of the field sites, only collar and root lesions were observed. However, at two sites, large stem and branch lesions unconnected to the collar region were also observed. Phytophthora lateralis was readily isolated from both aerial and basal lesions. In artificial inoculation experiments, two Scottish isolates of the pathogen caused lesions on C. lawsoniana shoots and were readily reisolated from the lesions, their pathogenicity being comparable to that of P. lateralis isolates originating from outside the UK. Isolates from six field sites and the two nursery interceptions exhibited ITS and cox II sequences identical to published sequences of French and North American isolates. However, the isolates from two field sites shared an ITS sequence with Taiwanese isolates and differed from North American, French and Taiwanese isolates by a single‐base substitution in cox II, suggesting a separate evolutionary history. It is clear that P. lateralis now presents a significant threat to C. lawsoniana in Britain. The main source of the outbreaks is likely to be imported infested nursery stock.     

Canker of cypresses caused by Coryneum cardinale Wag. in Britain

Coryneum canker occurs widely on Cupressus macrocarpa in England. Mycelial inoculations caused perennial cankers on Cupressocyparis leylandii and Chamaecyparis nootkatensis, severe but annual cankers on Thuja plicata and death of twigs on Juniperus communis. Chamaecy-paris lawsoniana rapidly overcame initial infection. Conidial inoculations on C. leylandii caused cankers. The potential of the disease in Britain and its control are discussed and aids to the identification of the fungus in culture given.     

Update on the 35‐year expansion of the invasive root pathogen,Phytophthora lateralis,across a landscape of Port Orford cedar (Chamaecyparis lawsoniana)

Port Orford cedar (Chamaecyparis lawsoniana) is endemic to northern California and southwestern Oregon and is considered a foundation species that plays critical roles in riparian areas and on nutrient‐poor soils. Since 1952, a non‐native, pathogenic oomycete (Phytophthora lateralis) has been spreading throughout the range of the cedar. Most spread occurs by vehicles carrying infested soil along gravel roads primarily used for timber harvest. In a previous study conducted in 1998 and 1999, Port Orford cedar and P. lateralis were censused in a 37‐km² study area and dendrochronology was used to reconstruct the history of pathogen invasion. That work, which represents the only detailed analysis of spread rates for P. lateralis, showed that the first successful invasion into the study area took place in 1977 and that 43% of the susceptible host sites (stream crossings) were infested by 1999. In the work presented here, all sites that were uninfested in 1999 were re‐censused in 2012, extending the historical reconstruction of P. lateralis spread to 35 years. Two new infestations were initiated between 1999 and 2012, suggesting that the rate of spread of P. lateralis has slowed greatly. Between 1980 and 1989, the average number of new site infestations was 1.8 infestations per year, while between 1990 and 1999 the average was 0.4 infestations per year and between 2000 and 2009 the average was 0.2 infestations per year. Several potential explanations for the reduced number of new infestations are discussed.     

Insights into the potential host range of Phytophthora foliorum

During a survey for Phytophthora ramorum undertaken in north‐west Scotland in early 2016, Phytophthora foliorum was found infecting foliage of the invasive shrub Rhododendron ponticum. Prior to this, P. foliorum had only been reported from foliage of hybrid azaleas in nurseries in California and Tennessee and from azalea plants in an ornamental nursery in Spain. No other hosts were known, and much of the behaviour of P. foliorum remained enigmatic. The species is classified in Phytophthora Clade 8c, with closest relatives, P. ramorum and Phytophthora lateralis, both of which are highly damaging tree pathogens. To explore the threat that P. foliorum might pose to trees, its growth–temperature responses on agar media and ability to cause lesions in the living bark of various hosts were contrasted with the behaviours of P. ramorum and P. lateralis. Phytophthora foliorum proved faster growing and more tolerant of temperature extremes than the other Phytophthora species. Comparisons of bark colonization initially focussed on R. ponticum and larch species Larix decidua and Larix kaempferi as all three are significant hosts of P. ramorum in the UK. Further experiments included another P. ramorum host, Fagus sylvatica (European beech), and the main host of P. lateralis, Chamaecyparis lawsoniana (Lawson cypress). Findings suggested that as well as being a significant pathogen of R. ponticum, damage caused by P. foliorum to both species of larch and beech was very similar to that of the EU1 lineage of P. ramorum, although growth in host tissue was also influenced by season.     

Root and butt rot caused by Heterobasidion annosum in Atlantic coniferous ecosystems of Spain

In the second half of the 20th century, the forested surface in northern Spain started to increase during a process of reforestation and the replacement of native forest. This reforestation was performed mostly by introducing monocultures of exotic coniferous species. One of the implications of intense forest exploitation and the introduction of new forest species is an increase in disease outbreaks. Because the genus Heterobasidion includes some of the most significant conifer pathogens in the world, surveys were conducted to collect and identify H eterobasidion isolates associated with diverse hosts within coniferous forests and plantations in the Basque Country, northern Spain. A total of 159 stands were surveyed, and 45 isolates were obtained from different trees. Based on sequencing of the internal transcribed spacer region of the ribosomal DNA, all collected isolates were identified as European H eterobasidion annosum s.s. (European P‐type). Heterobasidion annosum was detected in 28.3% of the sampled stands, with the following distribution by host: Chamaecyparis lawsoniana (11.1%), P inus pinaster (2.2%), P . radiata (42.2%), P . nigra (2.2%), P . sylvestris (17.8%), P seudotsuga menziesii (17.8%) and P icea abies (6.7%). The spatial distribution of the population showed a high degree of clustering. This is the first report of H . annosum s.s. causing damage to forest plantations of Chamaecyparis lawsoniana, P inus pinaster, P . radiata, P seudotsuga menziesii and P icea abies in Spain. In the current context of forest pathosystems, the management practice of replacing susceptible forest species with resistant species is recommended.     

Multilocus phylogenetic analysis revealed a new cryptic lineage of Serpula himantioides in Japan

Serpula himantioides is a widely distributed saprotrophic fungus that causes root and butt rot in various tree species. In Japan, butt rot associated with S. himantioides is observed in Chamaecyparis pisifera and Abies sachalinensis. Previous studies have shown that S. himantioides includes five phylogenetically defined cryptic lineages, but the placement of Japanese isolates remains unclear. To clarify the phylogenetic relationship between Japanese S. himantioides and the five known lineages, we conducted a phylogenetic analysis using three newly collected Japanese isolates along with 74 S. himantioides isolates, based on the internal transcribed spacer (ITS) region, the 28 S large subunit (LSU) rDNA, the beta-tubulin (tub) gene, and the heat stress protein (hsp) gene. The concatenated phylogenetic tree showed that Japanese isolates composed a clade with a high bootstrap value distinct from the known lineages, indicating the Japanese isolates present a new cryptic lineage of S. himantioides.     

Responses of Cupressus and Chamaecyparis callus tissues to inoculations with Seiridium cardinale

Inoculations of callus generated from micropropagated shoots of Cupressus sempervirens, Cupressus macrocarpa, Cupressus torulosa and Chamaecyparis lawsoniana showed that callus of species resistant or relatively resistant to Seiridium cardinale supported significantly less surface growth of the pathogen than callus of susceptible species. In dual cultures of Cupressus sempervirens or Cupressus torulosa callus with S. cardinale, inhibition of fungal growth towards the callus was correlated with known field resistance of these host species. Lignification, accumulation of polyphenolics and possible suberization of cell walls were detected in inoculated callus. This response was also correlated with known field resistance of the tested host species.     

Host specificity of co‐infecting Botryosphaeriaceae on ornamental and forest trees in the Western Balkans

The Botryosphaeriaceae is a diverse family of endophytes and fungal pathogens of mainly woody plants. We considered the host range and distribution of these fungi by sampling diseased ornamental and forest trees and shrubs in Serbia, Montenegro, Bosnia and Herzegovina, spanning a Mediterranean and a Continental climatic region. In total, ten Botryosphaeriaceae species were identified in the Western Balkans and with the exception of Sphaeropsis visci and Phaeobotryon cupressi, which occurred on one host, all the species had a broader host range. Phaeobotryon cupressi was found only in the Mediterranean region and S. visci, Dothiorella sp., Dothiorella sarmentorum and Diplodia seriata were present only in the Continental region. Pathogenicity tests were conducted on a variety of hosts from which the Botryosphaeriaceae species were isolated. These included leaves and/or stems of seedlings of 21 hosts, and cut leaves and/or branches of six hosts. Moreover, stems of seedlings of Chamaecyparis lawsoniana, Cedrus deodara, Picea omorika, Pinus patula and Eucalyptus grandis were inoculated as hosts from which some or all of the Botryosphaeriaceae species used for inoculation were not isolated. Inoculations showed that the majority of these fungi could also co‐infect hosts other than those from which they were isolated. The results suggest that most of the species have broad host ranges and can potentially cause disease on a broad range of tree species under certain conditions.     

Root and aerial infections of Chamaecyparis lawsoniana by Phytophthora lateralis: a new threat for European countries

Phytophthora lateralis has been isolated from root and collar lesions in Port‐Orford Cedar (POC) trees (Chamaecyparis lawsoniana) in north‐western France (Brittany). These trees, planted in hedgerows, displayed symptoms similar to the typical symptoms of POC root disease. Until now, the disease has been found outside of the nurseries only in western North America. Aerial symptoms, not associated with root or collar infections, were also observed, and P. lateralis was isolated from branch lesions. Similar symptoms were previously observed only in one POC root disease site, located in the Pacific coast of Oregon where climatic conditions are similar to those occurring in Brittany. The reported aetiology as well as the morphological characteristics (deciduous sporangia) of P. lateralis suggests that this species could be air‐dispersed, as described for P. ramorum, a closely related species. This outbreak of P. lateralis in Brittany in farming landscapes associated with the aerial spread of this pathogen represents a new threat for European countries.     

Colonisation of native and exotic conifers by indigenous bark beetles (Coleoptera: Scolytinae) in France

Planting exotic conifers offers indigenous forest insects an opportunity to extend their host range and eventually to become significant pests. Knowing the ecological and evolutionary modalities driving the colonisation of exotic tree species by indigenous insects is thus of primary importance. We compared the bark beetle communities (Coleoptera: Curculionidae, Scolytinae) associated with both native and introduced conifers in France. The aim of our study was to estimate the influence of both host- and insect-related factors on the beetles’ likelihood to shift onto new hosts. We considered the influence of host origin (i.e. native vs. exotic), host tree species identity, tree bark thickness and tree taxonomic proximity, as well as insects’ host specificity. A field inventory using trap trees was carried out in two regions in France (Limousin and Jura) during two consecutive years (2006 and 2007) on three European native conifer species [Norway spruce (Picea abies); Scots pine (Pinus sylvestris) and European Silver-fir (Abies alba)] and five North American [Sitka spruce (Picea sitchensis); Eastern white pine (Pinus strobus); Grand fir (Abies grandis); Douglas fir (Pseudotsuga menziesii) and Western red cedar (Thuja plicata)]. A total of 18 indigenous and 2 exotic bark beetle species were collected. All exotic conifer species were colonised by indigenous bark beetle species and no significant difference was observed of the cumulated species richness of the latter between native and exotic tree species (13 vs. 14, P < 0.05). The ability of indigenous bark beetles to shift onto exotic conifers appeared to strongly depend on host species (significantly structuring bark beetle assemblages), the presence of phylogenetically related native conifer species and that of similar resources, in combination with insect host specificity. Host tree species status (native or exotic) also seemed to be involved, but its effect did not seem as essential as that of the previous factors. These findings are discussed in terms of adaptation, plasticity and practical aspects of forest management.     

Isolation of Phytophthora lateralis from Chamaecyparis foliage in Taiwan

Following the discovery in 2008 of Phytophthora lateralis in forest soil under old‐growth yellow cedar (Chamaecyparis obtusa var. formosana) in north‐east Taiwan, further sampling was undertaken in the same region. Soil, root and symptomatic foliage samples were collected from five separate sites where C. obtusa was the dominant species in cloud forests at ca. 1800–2500 m. Soil and fine root samples were baited with cedar needles; both direct isolation and cedar needle baiting were used on foliage samples. Phytophthora lateralis was obtained from soil at three of the sites, but only from three of the 27 soil samples overall. Only one of 25 root samples yielded the pathogen, and this was associated with infested soil. Three foliage samples with symptoms visible as dark brown to black frond tips also yielded P. lateralis; these came from two different sites. This is the first record of P. lateralis infecting the foliage of C. obtusa. Moreover, when some of the symptomatic Chamaecyparis foliage segments were incubated, sporangia of P. lateralis formed on the necrotic tissues, sometimes in the axils of needle segments. The study provides evidence that P. lateralis has both a soil/root infecting phase and an aerial or foliar infecting phase in Taiwan, which is consistent with its unusual combination of water‐dispersed (non‐papillate) and aerially dispersed (caducous) sporangia. It also demonstrates the importance of investigating the biology, aetiology and ecological behaviour of Phytophthoras in their native, endemic environments.     

Port‐Orford‐cedar resistant to Phytophthora lateralis

Phytophthora lateralis, an exotic root pathogen, is the primary cause of Port‐Orford‐cedar (Chamaecyparis lawsoniana) mortality throughout its native range in southwest Oregon and northwest California. Most trees in the field are very susceptible, but genetic resistance to this pathogen has been demonstrated. Since the late 1980s, the USDA Forest Service and the Bureau of Land Management in cooperation with Oregon State University have conducted an intensive programme to identify and test resistant trees from the field, and propagate them in a seed orchard with the goal of providing resistant seedlings for regeneration. Susceptible families showed only 0–10% survival using a variety of inoculation techniques. Rooted cuttings of resistant parents are seldom killed, and seedling families of these parents exhibit 25–100% survival, depending on family and inoculation technique. Symptom development on resistant trees, including sunken lesions, and resinosis as well as reduced colonization and re‐isolation success, was consistent with a hypersensitive reaction. In a long‐term field test, five resistant families had 20–80% survival after 16 years, while three susceptible families had 0–8% survival in the same interval.     

Effects of heartwood extractive fractions of Thuja plicata and Chamaecyparis nootkatensis on wood degradation by termites or fungi

The effect of selective removal of extractives on termite or decay resistance was assessed with matched samples of Thuja plicata Donn ex D. Don and Chamaecyparis nootkatensis (D.Don) Spach heartwood. Samples were extracted using a variety of solvents and then exposed to the subterranean termite Coptotermes formosanus Shiraki in a no-choice feeding test or to the brown-rot fungus Postia placenta (Fr.) M. Larsen & Lombard in a soil bottle test. At the same time, the effect of naturally occurring variations in heartwood extractives on termite or decay resistance was evaluated by testing samples from the inner and outer heartwood of five trees of each species against C. formosanus and P. placenta and analyzing matched wood samples for their extractive content. The results suggest that the methanol-soluble extractives in T. plicata and C. nootkatensis play an important role in heartwood resistance to attack by C. formosanus and P. placenta. Total methanol-soluble extractive content of the heartwood was positively correlated with both termite and decay resistance; however, there was much unexplained variation and levels of individual extractive components were only weakly correlated with one another. Further studies are under way to develop a better understanding of the relationships between individual extractive levels and performance.     

Two conifer species native to Patagonia threatened by Phytophthora austrocedri

Phytophthora austrocedri is a pathogen of Austrocedrus chilensis causing “Mal del Ciprés” (cypress sickness) in Patagonia and killing Juniperus communis in Britain, along with other species in the Cupressaceae. The evident association of the pathogen with members of the Cupressaceae makes it necessary to study the susceptibility of other species related to A. chilensis to prevent the emergence of new diseases. The present study tested the pathogenicity of P. austrocedri to three native and endemic conifers in South America: Fitzroya cupressoides, Pilgerodendron uviferum and Araucaria araucana. The results showed that F. cupressoides and P. uviferum were highly susceptible to P. austrocedri. Since these are endangered tree species, the outcome of this study is crucial for governmental agencies, policymakers and stakeholders.     

Pathogenicity of Malaysian Phytophthora palmivora on cocoa,durian, rubber and oil palm determines the threat of bud rot disease

The epidemic of bud rot disease affecting oil palm in Colombia is primarily caused by Phytophthora palmivora. The pathogen has a cosmopolitan presence that includes Southeast Asia, but to date, bud rot has not been reported in this region. This study provides an overview of the potential risk of Malaysian P. palmivora isolates cross‐infecting other host species, including cocoa, durian, rubber and Malaysian oil palm planting materials (Dura × Pisifera, D × P). On cocoa pods, the durian isolate PP7 caused dark brown necrotic lesions. Detached leaf bioassays showed that P. palmivora isolates PP3 and PP7 infected different hosts, except rubber foliage without wounding. Inoculation tests on cocoa, durian and rubber seedlings caused brown necrotic lesions when stems were wounded, with 10% mortality in cocoa and durian at 17 days post‐inoculation (dpi). However, no further infection was observed, and lesions closed within 14–28 dpi on the non‐wounded seedlings. Pathogenicity tests of oil palm seedlings inoculated with isolates PP3 and PP7 indicated that Malaysian P. palmivora isolates were not pathogenic to oil palms based on localized infection observed only through wounding. Overall, the work demonstrated that Malaysian P. palmivora isolates were able to cross‐infect multiple hosts but did not show severe infections on oil palms.     

First report of Phyllosticta spinarum causing arborvitae foliar blight in Russia

Thuja occidentalis is a popular and widely planted tree in Russia. However, it is susceptible to arborvitae foliar blight, which can compromise the long-term health of trees in the landscape and in nursery production. Leaves with arborvitae needle blight were collected in 2019 in the foothill zone of the Republic of Adygea, and two isolates of Phyllosticta sp. were obtained. Multilocus analyses were performed with sequences of the internal transcribed spacer and 28 S rDNA of the ribosomal DNA, partial actin gene, partial RNA-polymerase II gene, and partial translation elongation factor 1-alpha. Based on phylogenetic analyses combined with morphological features the isolates were identified as Phyllosticta spinarum. Following inoculation of the original host species both isolates produced symptoms similar to those observed in naturally infected leaves. Koch's postulates were then fulfilled by re-isolating P. spinarum from the inoculated leaves. These findings represent the first confirmed detection of P. spinarum causing arborvitae needle blight in Russia.     

Ecology of Armillaria species on conifers in Japan

Distribution, host preference and pathogenicity of Japanese Armillaria species on conifers were investigated on the basis of field collections of 65 isolates. We identified seven Armillaria species from 19 conifer species including six major Japanese plantation conifers using mating tests and sequences of the translation elongation‐1 α gene. Armillaria mellea, Armillaria ostoyae, Armillaria cepistipes and Armillaria sinapina were frequently collected, whereas Armillaria nabsnona, Armillaria tabescens and a biological species Nagasawa’s E were rare. On the basis of host condition when the isolates were collected, A. mellea, A. ostoyae, A. cepistipes and A. tabescens are considered as moderate to aggressive pathogens of conifers in Japan.     

Agrobacterium pusense,a new plant tumour‐inducing pathogen isolated from Lawson cypress

Lawson cypress (Chamaecyparis lawsoniana), an important landscape tree, is widely planted in gardens and parks throughout Iran. Crown gall disease on Lawson cypress trees was observed in Sari and Juybar Counties, Mazandaran province, northern Iran, in 2017. Isolation from galls on potato dextrose agar (PDA) containing CaCO₃ yielded bacterial colonies, the predominant types of which were purified and selected for characterization. The isolates were Gram‐negative, oxidase positive, able to grow in 2% NaCl and produced 3‐ketolactose. They hydrolysed esculin, casein and arbutin but not starch, gelatin or Tween 80. Two representative isolates were selected for PCR amplification and sequencing of DNA gyrase subunit B (gyrB) gene. In the phylogenetic tree based on the partial sequence of the gyrB gene, isolates KH1 and KH2 clustered with Agrobacterium pusense. The pathogenicity of all isolates was confirmed by inoculation on Jimsonweed (Datura stramonium) and carrot discs (Daucus carota). Confirmation of the presence of genes involved in pathogenicity was made by performing PCR with the virD2A/virD2C and VCF/VCR primer pairs which resulted in amplification of the expected 224 and 730 bp fragments in all studied isolates, respectively. A. pusense was therefore identified as the causal agent of crown and stem gall of Lawson cypress. This appears to be the first report on the natural occurrence of crown gall disease on Lawson cypress and the first record of a plant disease caused by A. pusense.     

Phytophthora ramorum is a generalist plant pathogen with differences in virulence between isolates from infectious and dead‐end hosts

Variation in virulence was examined among isolates of Phytophthora ramorum from epidemiologically important or infectious (non‐oak) and transmissive dead‐end (oak) hosts from North America. Twelve isolates representative of the genetic, geographic and host range of P. ramorum in the western United States were inoculated on leaves of Umbellularia californica (bay laurel or bay) and stems of Quercus agrifolia (coast live oak). In spite of extreme genetic similarity among the isolates employed, and even within the same genotype, significant differences in lesion size were measured, suggesting virulence in this pathogen is also controlled by epigenetic factors. A strong positive correlation between lesion size on bay laurel and coast live oak provides experimental evidence P. ramorum is a generalist pathogen that lacks host specificity. Isolates from non‐transmissive oaks were significantly less pathogenic both on oaks and bays than isolates from infectious hosts. These results are essential to further our understanding of the epidemiology and evolutionary potential of this pathogen. A quantitative differential in virulence of isolates from hosts with different epidemiological roles has been described for many animal diseases, but is a novel report for a plant disease.     

Intraspecies variation of mature Pinus taeda in response to root‐infecting ophiostomatoid fungi

Seedling screening studies have shown intraspecies variation in susceptibility of Pinus taeda (loblolly pine) to Leptographium terebrantis and Grosmannia huntii, the causal agents of root infection in Pinus species. However, it is critical to understand the susceptibility of mature P. taeda trees. Roots of mature P. taeda families determined as susceptible and tolerant to L. terebrantis and G. huntii by previous seedling screening trials were artificially inoculated with the same fungal isolates. Dark necrotic lesion and vascular occlusion were recorded 8 weeks later. Families previously considered as susceptible had relatively longer lesions and occlusions. The variation in susceptibility to the two fungi remained the same as exhibited by families in the seedling trial. These results suggest intraspecies variation in relative susceptibility of P. taeda to L. terebrantis and G. huntii remains similar regardless of the age of the tree.